Carrot, a biennial herb of the Apiaceae family, is among the most important vegetable crops in the world. In this study, nine candidate reference genes (GAPDH, ACTIN, eIF-4α, PP2A, SAND, TIP41, UBQ, EF-1α, and TUB) were cloned from carrot. Carrot plants were subjected to abiotic stresses (heat, cold, salt, and drought) and hormone stimuli (gibberellin, salicylic acid, methyl jasmonate, and abscisic acid). The expression profiles of the candidate reference genes were evaluated in three technical and biological replicates. Real-time qPCR data analyses were performed using three commonly used Excel-based applets namely, BestKeeper, geNorm, and NormFinder. ACTIN and TUB were the most stable genes identified among all sample groups, but individual analysis revealed changes in their expression profiles. GAPDH displayed the maximum stability for most of single stresses. To further validate the suitability of the reference genes identified in this study, the expression profile of DcDREB-A1 gene (homolog of AtDREB-A1 gene of Arabidophsis) was studied in carrot. The appropriate reference genes were selected that showed stable expression under the different experimental conditions.

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Selection of suitable reference genes for qPCR normalization under abiotic stresses and hormone stimuli in carrot leaves.

Cold acclimation of winter cereals and other winter hardy species is a prerequisite to increase subsequent freezing tolerance. Low temperatures upregulate the expression of C-repeat/dehydration-responsive element binding transcription factors (CBF/DREB1) which in turn induce the expression of COLD-REGULATED (COR) genes. We summarize evidence which indicates that the integration of these interactions is responsible for the dwarf phenotype and enhanced photosynthetic performance associated with cold-acclimated and CBF-overexpressing plants. Plants overexpressing CBFs but grown at warm temperatures mimic the cold-tolerant, dwarf, compact phenotype; increased photosynthetic performance; and biomass accumulation typically associated with cold-acclimated plants. In this review, we propose a model whereby the cold acclimation signal is perceived by plants through an integration of low temperature and changes in light intensity, as well as changes in light quality. Such integration leads to the activation of the CBF-regulon and subsequent upregulation of COR gene and GA 2-oxidase (GA2ox) expression which results in a dwarf phenotype coupled with increased freezing tolerance and enhanced photosynthetic performance. We conclude that, due to their photoautotrophic nature, plants do not rely on a single low temperature sensor, but integrate changes in light intensity, light quality, and membrane viscosity in order to establish the cold-acclimated state. CBFs appear to act as master regulators of these interconnecting sensing/signaling pathways.

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Role of CBFs as Integrators of Chloroplast Redox, Phytochrome and Plant Hormone Signaling during Cold Acclimation

To farmers, horticulturalists, and others whose livelihood depends on growing plants, it is obvious that the transition from vegetative to reproductive development is a critical phase in the life cycle of higher plants. It is difficult to overestimate the impact the direct products of flowering have on human endeavors. Because they are an integral part of the human diet, production of seeds and fruits form the foundation of all nations’ economies. Thus, it becomes equally evident that the ability to manipulate and control flowering with simple treatments has enormous potential both from an economic standpoint as well as increasing food production for an ever growing human population. However, the development of such cultural techniques is predicated on a thorough understanding of the physiological, biochemical, and molecular aspects of reproductive development. Although a great deal of descriptive information exists for many species about the influence of environmental factors on reproductive development, knowledge of the mechanisms by which the floral transition takes place is almost totally lacking. Flowering, or more precisely, reproductive development, is composed of many independent but highly coordinated processes. To discuss the role of hormones in floweringper sewould be pointless; it is much more convenient to arbitrarily divide flowering into several temporally related sequences.

Hormones and Reproductive Development

Low temperature is one of the important limiting factors for growing season and geographical distribution of plants. Zoysiagrass (Zoysia Willd) is one of the widely used warm-season turfgrass that is distribute in many parts of the world. Zoysaigrass native to high latitude may have evolved higher cold tolerance than the ones native to low latitude. The objective of this study was to investigate the cold stress response in zoysiagrass native to diverse latitude at phenotypic, physiological and metabolic levels. Two zoysiagrass (Z. japonica) genotypes, Latitude-40 (higher latitude) and Latitude-22 (lower latitude) were subjected to four temperature treatments (optimum, 30/25°C, day/night; suboptimum, 18/12°C; chilling, 8/2°C; freezing, 2/-4°C) progressively in growth chambers. Low temperature (chilling and freezing) increased leaf electrolyte leakage (EL) and reduced plant growth, turf quality, chlorophyll (Chl) content, photochemical efficiency (Fv/Fm) and photosynthesis (Pn, net photosynthetic rate; gs, stomatal conductance; intercellular CO2; Tr, transpiration rate) in two genotypes, with more rapid changes in Latitude-22. Leaf carbohydrates content (glucose, fructose, sucrose, trehalose, fructan, starch) increased with the decreasing of temperature, to a great extend in Latitude-40. Leaf abscisic acid (ABA), salicylic acid (SA) and jasmonic acid (JA) content increased, while indole-3-acetic acid (IAA), gibberellic acid (GA3) and trans-zeatin ribside (t-ZR) content decreased with the reduction of temperature, with higher content in Latitude-40 than in Latitude-22. Chilling and freezing induced the up-regulation of C-repeat binding factor (ZjCBF), late embryogenesis abundant (ZjLEA3) and dehydration-responsive element binding (ZjDREB1) transcription factors in two genotypes, whereas those genes exhibited higher expression levels in Latitude-40, particularly under freezing temperature. These results suggested that zoysiagrass native to higher latitude exhibited higher freezing tolerance may attribute to the higher carbohydrates serving as energy reserves and stress protectants that stabilize cellular membranes. The phytohormones may serve signals in regulating plant growth, development and adaptation to low temperature as well as inducing the up-regulated ZjCBF, ZjLEA3 and ZjDREB1 expressions thus result in a higher cold tolerance.

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Differential physiological and metabolic response to low temperature in two zoysiagrass genotypes native to high and low latitude

This chapter is intended to provide an overview of the functions of the five major plant growth hormones in the intact plant: auxin (IAA), gibberellins (GA), cytokinins (K), ethylene, and abscisic acid (ABA). The effects of these hormones have been described in detail on numerous occasions over the last 10 years (see general references at the end of this chapter). This review utilizes these previous reviews as well as including a survey of the literature published between January 1977 and January 1980. The most recent and surely the most detailed treatment of the subject is to be found in Phytohormones and Related Compounds: A Comprehensive Treatise. D.S. Letham et al., Volumes 1 and 2, 1978. The intent of the present review is to provide a more general overview and to consider overall patterns of regulatory controls by hormones in the whole plant.

Functions of Hormones at the Whole Plant Level of Organization

Stecklings (roots) of three cultivars of carrots (Daucus carota L.) were vernalized 10 weeks at 5 C and subsequently grown at each of three greenhouse night/day temperature regimes: high (27/32 C), medium (21/27 C), and low (15/21 C). Floral differentiation occurred first in the easy bolting cv. Scarlet Nantes, intermediate in cv. Danvers 126, and last in cv. Royal Chantenay. Stem elongation arising from the subapical meristematic region always preceded floral differentiation. Extractable gibberellin-like activity in carrot stem apices increased from harvest during the 10-week vernalization period, then remained constant even though floral differentiation and stem elongation occurred during an additional 20-week cold storage period. Low temperature had both an inductive and a direct effect on reproductive development depending on length of low temperature exposure.After 10 weeks vernalization at 5 C, high greenhouse temperature severely reduced ultimate seedstalk height and the endogenous gibberellinlike activity decreased rapidly during the first 3 weeks in the greenhouse. At the low greenhouse temperature, activity remained fairly constant during the 10-week sampling period. Changes in endogenous gibberellinlike activity were related with stem elongation, but not with floral initiation. Exogenous gibberellic acid (GA(3)) applied following vernalization prevented the inhibitory effect of high greenhouse temperature on seedstalk elongation and resulted in seedstalk heights comparable to untreated controls grown at the low greenhouse temperature. Exogenous applications of succinic acid-2,2-dimethylhydrazide and chlormequat reduced seedstalk height of carrot plants grown at the medium and low greenhouse temperatures to that of untreated controls grown at high temperature. Exogenous growth regulators and greenhouse temperature affected seedstalk elongation, but did not affect the number of plants that flowered.

William C. Kelly

Papers

Temperature Interactions with Growth Regulators and Endogenous Gibberellin-like Activity during Seedstalk Elongation in Carrots.